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Zomation and Differentiation of Tissues in the Primary ZOMATION AND DIFFERENTIATION OF TISSUES IN THE PRIMARY ROOT OF SOYBEAN DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University 3 y CHAO NIEN SUN, B. Sc., M. Sc. The Ohio State University 1953 Approved by: Adviser {/ ACKNOWLEDGEMENT This investigation was carried out under the direction of Dr. R. A. Popham of the Department of Botany and Plant Pathology, The Ohio State University, Columbus, Ohio. The writer wishes to express his deep gratitude to him and also to all who have aided in any way during the course of this study and in the preparation of this paper. i A Q 9 8 4 3 TABLE OF CONTENTS I. ORGANIZATION OF THE ROOT APICAL MERISTEM INTRODUCTION ............................................... 1 MATERIALS AND METHODS ......... ............................ k GENERAL PATTERN OF ZONATION ................................ 7 1. The stelar initials and their derivatives . ....... 7 2. The common initials and their derivatives ........ 8 3* Comparison of apices of primary roots of various ages ...... 9 DISCUSSION ................................................. 16 SUMMARY ..................................................... 18 LITERATURE CITED ........................................... 19 II. GROWTH AND TISSUE DIFFERENTIATION IN PRIMARY ROOTS ........ 21 PROCEDURES .... 22 EXPERIMENTAL RESULTS ....................................... 23 1. External morphology.............................. 23 2. General structure of the primary root ............ 25 3- Tissue differentiation........ 27 A. Primary phloem ................. 27 B. Primary xylem ................................ 28 C. Stelar' cambium .......................... 3h D . Pericycle ......................... R2 E. Pericycle - secondary root origin ....... 1x3 F. Endodermis ................................... U3 G. Cortical parenchyma .......................... AA H. Epidermis and root hairs ..................... ilA ii I. Secondary tissues ....................... 1+5 J. Root cap ..................................... 1+5 K. Comparison of growth region and tissue differentiation in aerated and non-aerated roots ........................................ 1+6 DISCUSSION ............................................ 1+7 SUMMARY.................................................... h9 LITERATURE CITED .................. 51 AUTOBIOGRAPHY ................................................... 53 iii ZONATION AND DIFFERENTIATION OF TISSUES IN THE PRIMARY ROOT OF SOYBEAN I. ORGANIZATION OF THE ROOT APICAL MERISTEM INTRODUCTION Much work has been done within the last decade on zonation structure of shoot apices, but recent investigations on root apical meristems are surprisingly few. The present study undertakes to clarify the details of zonation in the soybean root tip. Perhaps the earliest study of the root apex is that by NM-geli (18145)* Some 25 years later, Hanstein (1870) founded the well known histogen theory of apical meristem organization. Although his concept is not applicable to shoot meristems, it is still useful in describing the organization and origin of root tissues. Later workers, such as * Janczewski (l87li)> Eriksson (1878), and Flahaut (1878), classified root meristems according to their interpretation of the histogen theory. Because soybean belongs to Leguminosae, a brief outline of the structural differences among root apices of Leguminosae studied and recorded to date will be presented. Janczewski (I87I4) described five types of promeristems for the phanerogams. In his fourth type, for which the roots of some Leguminosae (Pisurn sativum and Phaseplus vulgaris) furnish examples, all tissues (the central cylinder, cortex, epidermis, and root cap) originate from a common meristematic zone. Eriksson (I878) distinguished four types of root apices for dicotyledons. His third type corresponds to Janczewski*s fourth type, being characterized by the fact that all of the primary tissues of the root originate from a common meristematic zone. In this class he places 2 the following species of Leguminosae: Vicia sativa, V. narbonensis, Pisum sativum, Cicer arietinum, Phaseplus multiflorus, Lathyrus odoratus, L. latifolius, Robinia pseudacacia, and Cassia glauca. Eriksson's fourth type, on the other hand, is characterized by the presence, at the apex of the root, of two different meristems, a plerome and a group of common initials. The former gives rise to the central cylinder. The latter gives rise to the cortex, epidermis and root cap. The root cap consists of two sharply defined parts: the columella and the peripheral portion. The columella is derived from common initials by transverse cell divisions while the peripheral part of the root cap is derived from the common initials by tangential cell divisions. In this group he placed Lupinus nanus, L. mutalilis, L. hybridus, L. albus, L. grandiflorus, L. dunetti, Mimosa pudica, and Acacia lophanta. Flahault (1878) examined individuals of three hundred and fifty species of Phanerogams and found that those examined in the genera Lupinus, Cercis, Gymnocladus, Guilandinia, Acacia, and Mimosa had distinct stelar initials and a common group of initials from which developed cortex, epidermis, and root cap. Tiegs (1912) x*eported three histogens, plerome, periblem, and protoderm-columella, in the promeristem of roots of three species of Leguminosae (Vicia villosa - lateral roots, Pisum sativum, and Trefolium repens). The central cylinder originates from the plerome, the cortex from the periblem, and the epidermis as well as the root cap from the protoderm-columella initials. Schliepp (1926), summarizing information on root meristems, 3 distinguished two zones in the root apexs corpus and tunica. The difference between corpus and tunica results from the pattern of division of their cells. By his definition, that part of the root where the number of cell rows increase away from initial zone is called corpus, while the number of cell rows increase towards the initial zone is called tunica. Since Schiiepp’s study dealt only with the pattern of derivation of cells, it is not possible, to correlate it directly with Hanstain's histogen theory. ‘Neuman (1939) found that in Mimosa and Lupinus the plerome and periblem originate from a central cell. Hie central cylinder is derived from the plerome, the cortex from the periblem. The columella of the root cap originates from columella initials. The peripheral part of the root cap originates by periclinal divisions in the "dermatogen" according to Neuman. The only investigation on zonation in the soybean (variety Mammoth Xellow) root tip which has come to the author's attention is that of Bell (193U)- Be found that the pattern of development corresponded to that of Janczewski's fourth class of angiosperms, namely, that the stele, cortex, epidermis, and root cap, instead of arising from definite histogens, originate from a common group of meristematic cells. The results of the present study, however, indicate a more complex root apex organization in the soybean. MATERIALS AND METHODS The primary roots of the Monroe variety of Glycine max ,L. Merrill were used in this study. The plants were cultured in an aerated four-salt solution under controlled illumination and temperature. Soybean seeds were first soaked for 2k hours in the culture solution. Then* in order to reduce or to eliminate surface contamination* they were treated for five minutes in a dilute disinfectant consisting of one part chlorox (5-25 per cent sodium hypochlorite by weight) to 20 parts water* a treatment which considerably increased the percentage of seeds germinating. The seeds were then placed in ’’germination crocks.u The l|r liter crocks were covered with cotton mosquito netting to support the beans and the culture solution level was maintained slightly above the cotton netting in order to keep the seeds moist. After 5 days* seedlings, selected on the basis of uniformity of size and length of primary roots, were transferred from the germination crocks to quart mason jars. The jars were painted black on the outside in order to avoid the growth of algae in the culture solution. One seedling was placed in each jar, the root being inserted through a hole in the rubber stopper. Aerators for both germination crocks and jars were made of £ cm. pieces of porous carbon tubing obtained from National Carbon Co.* Cleveland, Ohio. One end of the carbon tube was plugged with a rubber stopper* the other end was attached to an 18 cm. length of glass tubing. These aerators were then connected in parallel to an air line by means of 11T11 tubes. In the germination crocks, the aerator tubes were 5 inserted through the mosquito netting. In the jars, the aerator tubes were inserted through a hole in the rubber stopper used to support the seedlings. The plants were cultured in a ventilated room under a bank of twelve 96-inch General Electric, T-8, kf?00 white fluorescent tubes. Eighteen 60-watt incandescent bulbs, nine on each side of the bank of fluorescent tubes, were set up to provide supplementary light in the red end of the spectrum. The bank of lights was adjusted at the top of the plants to give a light intensity of 900-1000 foot candles as measured with Weston Illumination Meter - model 756, and was adjusted occasionally to compensate for lamp ageing. A photoperiod of 15 hours was maintained throughout the experiment. The temperature of the ventilated room was kept at 16-18° C. from 8 P. M. to 5 A. M. (dark hours) and at 22-2h° C. from 5 A. M. to
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